Feeding and egg distribution studies of Heringia calcarata (Diptera: Syrphidae), a specialized predator of woolly apple aphid (Homoptera: Eriosomatidae) in Virginia apple orchards

Predation by the aphidophagous syrphid fly Heringia calcarata (Loew) on woolly apple aphid, Eriosoma lanigerum (Hausmann), was studied in the laboratory and in Virginia apple orchards. Feeding studies compared the prey suitability of three temporally sympatric aphid pests of apple: spirea aphid, Aphis spiraecola Patch; rosy apple aphid, Dysaphis plantaginea (Passerini); and woolly apple aphid. Significantly more H. calcarata larvae survived and completed development on a pure diet of woolly apple aphid than on rosy apple aphid, and none survived on spirea aphid. Final larval weights were significantly greater, and the larval developmental period was significantly shorter on woolly apple aphid than on rosy apple aphid, but neither the duration of pupal development nor adult weight differed between diets. H. calcarata larvae consumed an average of 105 woolly apple aphids during their development. Na?ve, neonate larvae given access to all possible pair combinations of woolly apple aphid, rosy apple aphid, and spirea aphid consumed significantly more woolly apple aphids in all pairings that included woolly apple aphid. When given a choice of rosy apple aphid and spirea aphid, significantly more rosy apple aphids were consumed. Weekly counts of syrphid eggs found in woolly apple aphid, rosy apple aphid, and spirea aphid colonies collected from apple trees showed that two generalist hover fly predators, Eupeodes americanus (Wiedemann) and Syrphus rectus Osten Sacken, were present in colonies of all three aphid species and that E. americanus was the most abundant syrphid predator in A. spiraecola and D. plantaginea colonies. H. calcarata eggs were found only in woolly apple aphid colonies and were more abundant there than E. americanus and S. rectus. These data suggest that H. calcarata is a specialized predator of woolly apple aphid in the apple ecosystem in Virginia.     

Conservation biological control of rosy apple aphid, Dysaphis plantaginea (Passerini), in eastern North America

Because of the potentially serious damage rosy apple aphid, Dysaphis plantaginea (Passerini) (Homoptera: Aphididae), can cause to apple fruit and branch development, prophylactic insecticides are often used for control. If biological control could be relied on, the amount of pesticide applied in orchards could be reduced. This study examined biological control of rosy apple aphid in eastern West Virginia and the potential for enhancement through conservation biological control, in particular, the effect of interplanting extrafloral nectar-bearing peach trees. By 20 d after first bloom, only 2% of fundatrices initially present survived to form colonies based on regression of data from 687 colonies. Exclusion studies showed that many of the early colonies were probably destroyed by predation; the major predator responsible seemed to be adult Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Mortality before apple bloom was most important in controlling rosy apple aphid population growth but by itself is not sufficiently reliable to prevent economic injury. Interplanting of extrafloral nectar-bearing trees did not increase biological control, and interplanting with 50% trees with extrafloral nectar glands reduced biological control. The number of leaf curl colonies in the 50% interplanted orchards was lower than in monoculture orchards, suggesting a preference of alate oviparae for more diverse habitats, supporting the resource concentration hypothesis but not at a level sufficient to prevent injury. Predation and parasitism after the formation of leaf curl colonies was not adequate to control rosy apple aphid populations.     

Flower strips increase the control of rosy apple aphids after parasitoid releases in an apple orchard

Mass releases of two parasitoid species, Aphidius matricariae and Ephedrus cerasicola, may provide an alternative measure to pesticides to control the rosy apple aphid Dysaphis plantaginea in organic apple orchards. As an exploratory study, we tested if the presence of flower strips between apple tree rows could improve the action of three early parasitoid releases––and of other naturally present aphid enemies––on the control of aphid colonies and the number of aphids per tree. Apple trees located at various distances from parasitoid release points were monitored in plots with and without flower strips in an organic apple orchard over two years, along the season of aphid infestation (March to July). Our case study demonstrated that the presence of flowering plant mixes in the alleyways of the apple orchard reduced the presence of D. plantaginea by 33.4%, compared to plots without flower strips, at the infestation peak date. We also showed a negative effect of increasing the distance to parasitoid release points on aphid control. However, our results at the infestation peak date suggest that the presence of flower strips could marginally compensate for the detrimental effect of increasing distance to the release point, probably by improving the persistence and dispersal capacities of natural enemies. Despite high variations in aphid population dynamics between years, we conclude that combining flower strips with early parasitoid releases in apple orchards is promising for biological control of the rosy apple aphid, although the method merits to be further refined.     

Estimating gene flow between refuges and crops: a case study of the biological control of Eriosoma lanigerum by Aphelinus mali in apple orchards

Parasitoid disturbance populations in agroecosystems can be maintained through the provision of habitat refuges with host resources. However, specialized herbivores that feed on different host plants have been shown to form host-specialized races. Parasitoids may subsequently specialize on these herbivore host races and therefore prefer parasitizing insects from the refuge, avoiding foraging on the crop. Evidence is therefore required that parasitoids are able to move between the refuge and the crop and that the refuge is a source of parasitoids, without being an important source of herbivore pests. A North-South transect trough the Chilean Central Valley was sampled, including apple orchards and surrounding Pyracantha coccinea (M. Roem) (Rosales: Rosacea) hedges that were host of Eriosoma lanigerum (Hemiptera: Aphididae), a globally important aphid pest of cultivated apples. At each orchard, aphid colonies were collected and taken back to the laboratory to sample the emerging hymenopteran parasitoid Aphelinus mali (Hymenoptera: Aphelinidae). Aphid and parasitoid individuals were genotyped using species-specific microsatellite loci and genetic variability was assessed. By studying genetic variation, natural geographic barriers of the aphid pest became evident and some evidence for incipient host-plant specialization was found. However, this had no effect on the population-genetic features of its most important parasitoid. In conclusion, the lack of genetic differentiation among the parasitoids suggests the existence of a single large and panmictic population, which could parasite aphids on apple orchards and on P. coccinea hedges. The latter could thus comprise a suitable and putative refuge for parasitoids, which could be used to increase the effectiveness of biological control. Moreover, the strong geographical differentiation of the aphid suggests local reinfestations occur mainly from other apple orchards with only low reinfestation from P. cocinnea hedges. Finally, we propose that the putative refuge could act as a source of parasitoids without being a major source of aphids.     

Genetic diversity of woolly apple aphid Eriosoma lanigerum (Hemiptera: Aphididae) populations in the Western Cape, South Africa

The woolly apple aphid Eriosoma lanigerum (Hausmann) is one of the most damaging apple pests in South Africa. Information on its genetic diversity is lacking and this study, in which the genetic structure of parthenogenetic E. lanigerum populations was characterized in the Western Cape Province of South Africa, represents the first local study of its kind. A total of 192 individuals from four different regions were collected and analysed using amplified fragment length polymorphism (AFLP). Using five selective AFLP primer pairs, 250 fragments were scored for analysis. Results indicated that a low level of genetic variation was apparent in E. lanigerum populations in the Western Cape (H = 0.0192). Furthermore, populations collected from geographically distant regions were very closely related, which can partly be explained by the fact that agricultural practices were responsible for dissemination of populations from a common ancestor to geographically distant areas. The low level of variation found indicated that the possibility of controlling E. lanigerum in the Western Cape using host plant resistance is favourable. This is the first report of AFLP being used to characterize the genetic structure of an aphid species. Results indicate that this marker may be useful for analysis of other aphid species.     

Temporal and spatial variability of rosy apple aphid Dysaphis plantaginea populations: is there a role of the alternate host plant Plantago major?

• 1 The rosy apple aphid Dysaphis plantaginea (Passerini) (Homoptera: Aphididae) is a pest of economic importance to the apple industry worldwide, particularly in organic apple orchards where no acceptable controls are available. In the Similkameen Valley of British Columbia, Canada, the rosy apple aphid population size varies widely between orchards and between years. To explain this variation, potential environmental correlates of aphid density were evaluated. The architecture of the alternate host was also evaluated for its effect on rosy apple aphid summer survival and reproduction.
• 2 The percentage of trees infested by rosy apple aphids among orchards was in the range 8–94% for trees having at least one cluster with more than ten aphids in 2007 and in the range 0–39% in 2008.
• 3 A general linear model correlating aphid densities to the environmental variables of abundance of the alternate host (plantain Plantago spp.), foliar nitrogen, tree age and planting density, and reduced by stepwise regression, indicated that foliar nitrogen and tree age explained 33% of the variation. Abundance of the summer, alternate food plant, plantain, was not related to later aphid densities on apple trees.
• 4 Plantain architecture, however, influenced aphid numbers and 25‐fold more aphids were found on low‐lying plantain leaves than on more upright leaves. Experimental manipulation of leaf angle and leaf size showed that significantly more aphids occurred on low angle, large leaves. Finally, mowing that encouraged low lying plants prior to spring aphid migration was associated with a four‐fold greater number of both winged and wingless aphids on the plantain.

     

Population genetic structure of codling moth (Lepidoptera: Tortricidae) from apple orchards in central Chile

Codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), is the main pest of pome fruits worldwide. Despite its economic importance, little is known about the genetic structure and patterns of dispersal at the local and regional scale, which are important aspects for establishing a control strategy for this pest. An analysis of genetic variability using microsatellites was performed for 11 codling moth populations in the two major apple (Malus domestica Borkh) cropping regions in central Chile. Despite the geographical distances between some populations (approximately 185 km), there was low genetic differentiation among populations (F(ST) = 0.002176), with only slight isolation by distance. Only approximately 0.2% of the genetic variability was found among the populations. Geographically structured genetic variation was independent of apple orchard management (production or abandoned). These results suggest a high genetic exchange of codling moth between orchards, possibly mediated by human activities related to fruit production.     

DL‐β‐Aminobutyric acid application negatively affects reproduction and larval development of the rosy apple aphid,Dysaphis plantaginea,on apple

The rosy apple aphid, Dysaphis plantaginea (Passerini) (Hemiptera: Aphididae), is one of the major pests of European apple orchards, commonly controlled by the use of synthetic insecticides. In the present work, the non‐protein amino acid DL‐β‐aminobutyric acid (BABA), known to induce plant resistance against a wide range of abiotic and biotic stresses, has been tested for its protective effect against this pest on apple. We first verified the lack of any contact effect of BABA on the insect itself. Next we applied BABA as a soil drench to apple and monitored its effect on the population development of aphids after artificial infestation. We demonstrated that BABA strongly reduced the population growth and that this compound severely affected various life‐history characteristics of the aphid such as female longevity and fecundity, nymph mortality, and larval development.     

QTL analysis for aphid resistance and growth traits in apple

The rosy apple aphid (Dysaphis plantaginea), the leaf-curling aphid (Dysaphis cf. devecta) and the green apple aphid (Aphis pomi) are widespread pest insects that reduce growth of leaves, fruits and shoots in apple (Malus × domestica). Aphid control in apple orchards is generally achieved by insecticides, but alternative management options like growing resistant cultivars are needed for a more sustainable integrated pest management (IPM). A linkage map available for a segregating F₁-cross of the apple cultivars ‘Fiesta’ and ‘Discovery’ was used to investigate the genetic basis of resistance to aphids. Aphid infestation and plant growth characteristics were repeatedly assessed for the same 160 apple genotypes in three different environments and 2 consecutive years. We identified amplified fragment length polymorphism (AFLP) markers linked to quantitative trait loci (QTLs) for resistance to D. plantaginea (‘Fiesta’ linkage group 17, locus 57.7, marker E33M35–0269; heritability: 28.3%), and to D. cf. devecta (‘Fiesta’ linkage group 7, locus 4.5, marker E32M39–0195; heritability: 50.2%). Interactions between aphid species, differences in climatic conditions and the spatial distribution of aphid infestation were identified as possible factors impeding the detection of QTLs. A pedigree analysis of simple sequence repeat (SSR) marker alleles closely associated with the QTL markers revealed the presence of the alleles in other apple cultivars with reported aphid resistance (‘Wagener’, ‘Cox’s Orange Pippin’), highlighting the genetic basis and also the potential for gene pyramiding of aphid resistance in apple. Finally, significant QTLs for shoot length and stem diameter were identified, while there was no relationship between aphid resistance and plant trait QTLs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Role of aphid predator guild in controlling spirea aphid populations on apple in West Virginia,USA

Spirea aphid populations and their predators were studied on apple to identify predators of importance in controlling aphid populations. Methods included random and non-random sampling from apple orchards in West Virginia, USA, sentinel aphid colonies, laboratory feeding studies, and predator exclusion studies. Aphidoletes aphidimyza (Diptera: Cecidomyiidae), chrysopids (Neuroptera: Chrysopidae), Harmonia axyridis (Coleoptera: Coccinellidae), and Orius insidiosus (Hemiptera: Anthocoridae) were the most abundant predators associated with spirea aphid colonies on apple. Parasitoids were all but absent in the study. Abundance of all predators was density dependent with greater responses to aphid populations at the orchard scale than to tree or individual colony scales. A. aphidimyza, O. insidiosus, chrysopids, and syrphids (Diptera) had the greatest degree of density dependence on aphid populations, and spiders showed inverse density dependence. Exclusion of predators with both cages and insecticides produced significantly higher aphid populations. Because of high abundance, good synchrony with aphid populations, and high impact per individual, H. axyridis adults were the most important spirea aphid predator on apple.     

Baseline susceptibilities to imidacloprid for green apple aphid and spirea aphid (Homoptera: Aphididae) collected from apple in the Pacific northwest

Susceptibilities to the neonicotinyl insecticide imidacloprid were determined for clones of apple aphid, Aphis pomi De Geer, and spirea aphid, Aphis spiraecola Patch, collected from conventional and organic apple orchards and from crab apple and wild apple in Washington state and British Columbia over a period of 6 yr. For aphids collected during 1996--1998, adults were dipped in test solutions by using the Food and Agriculture Organization protocol, and third instars and adults were reared on treated apple leaf disks. During the final 3 yr of study, bioassays involved only third instars on treated leaf material. Tests showed that A. spiraecola was significantly more tolerant to imidacloprid compared with A. pomi. Depending on the bioassay method and aphid developmental stage, average LC50 values for A. spiraecola were 4.4 -5.7 times higher than those for A. pomi established under the same test conditions. Clones of both species from Washington were marginally more tolerant to imidacloprid than clones from British Columbia, but the differences were generally not significant. Average measures of susceptibility for clones from organic orchards or unsprayed trees also did not differ from those for clones from conventional orchards, and there was no evidence for increasing LC50 values over the 6 yr of study. Differences in susceptibility to insecticides between these two anatomically similar species should be considered during the testing of new products for use on apple.     

Kaolin particle films suppress many apple pests, disrupt natural enemies and promote woolly apple aphid

Multiple applications of hydrophobic kaolin particle film in apple orchards suppressed numbers of blossom weevil (Anthonomus pomorum), brown leaf weevil (Phyllobius oblongus), attelabid weevil (Caenorhinus pauxillus), leafhoppers (Empoasca vitis and Zygina flammigera) and green apple aphid (Aphis pomi) colonies. The kaolin treatments reduced the apple sawfly (Hoplocampa testudinea) fruit infestation on cultivar J. Grieve, and the fruit damage caused by oyster scale (Quadraspidiotus ostreaeformis), mussel scale (Lepidosaphes ulmi), early caterpillars, leaf rolling moths (Tortricidae), fruitlet‐mining tortrix moth (Pammene rhediella) and codling moth (Cydia pomonella). There was no effect on the number of colonies of rosy leaf curling aphid (Dysaphis devecta), nor on the fruit damage caused by common earwig (Forficula auricularia) and apple sawfly on cv. G. Delicious. The level of infestation of rosy apple aphid (Dysaphis plantaginea), leaf miner moths (Phyllonorycter blancardella, Lyonetia clerkella), and agromyzid flies (Phytomyza heringiana) increased in the kaolin‐treated plots. Kaolin treatments promoted woolly apple aphid (Eriosoma lanigerum) infestation, which became severe, while it reduced the abundance of polyphagous predators like F. auricularia, predaceous Heteroptera and Coleoptera, the red velvet mite (Allothrombium fuliginosum), spiders (Araneae) and the abundance of common black ant (Lasius niger). The treatments also reduced parasitism of the apple sawfly by the ichneumonid Lathrolestes ensator. Many weeks after ending the kaolin treatments, the number of predaceous Coleoptera and especially the number of spiders remained low in the kaolin‐treated plots.     

The effect of ant attendance on the success of rosy apple aphid populations, natural enemy abundance and apple damage in orchards

1 The rosy apple aphid, Dysaphis plantaginea, is the most serious pest of apple in Europe and, although conventionally controlled by insecticides, alternative management measures are being sought. Colonies of D. plantaginea are commonly attended by ants, yet the effects of this relationship have received little attention. 2 An ant exclusion study was conducted in two distant orchards within the U.K. At both sites, ants were excluded from a subset of D. plantaginea infested trees at the beginning of the season and populations were monitored. The number of natural enemies observed on trees was also recorded and, before harvest, the percentage of apples damaged by D. plantaginea calculated. 3 Overall, the exclusion of ants reduced the growth and eventual size of D. plantaginea populations. On trees accessed by ants, greater numbers of natural enemies were recorded, presumably because aphid populations were often greater on such trees. However, this increased natural enemy presence was diluted by the larger aphid populations such that individual aphids on ant‐attended trees were subjected to a lower natural enemy pressure compared with those on ant‐excluded trees. 4 At harvest, apple trees that had been accessed by ants bore a greater proportion of apples damaged by D. plantaginea. There were also differences in cultivar susceptibility to D. plantaginea damage. 5 The present study highlights the importance of the ant–D. plantaginea relationship and it ia suggested that ant manipulation, whether physically or by semiochemicals that disrupt the relationship, should be considered as a more prominent component in the development of future integrated pest management strategies.     

Aphis pomi (Hemiptera: Aphididae) population development, shoot characteristics, and antibiosis resistance in different apple genotypes

In high-value crops such as apple, Malus X domestica (Borkh.), insecticidal pest control is of high relevance. The use of resistant apple cultivars can increase the sustainability of pest management in apple orchards. Besides variation in plant chemistry that may influence plant resistance by antibiosis or antixenosis, plant growth characteristics also can affect plant susceptibility to pests such as aphids. Variable susceptibility to the apple aphid, Aphis pomi De Geer (Hemiptera: Aphididae), has been described for different apple cultivars. These observations were based on phenotypic surveys and no information on genetically based apple resistance to A. pomi is yet available. The objective of this study was to relate shoot growth characteristics with aphid population development, and to assess the genetic background of apple antibiosis-based resistance to A. pomi by quantitative trait loci (QTL) analysis. Aphid population development was repeatedly studied in the field in sleeve cages attached to 200 apple trees of different genotypes. Aphid population development was positively correlated to shoot length and growth, and it also was affected by climatic conditions. Indications for antibiosis-based resistance to A. pomi remained weak in the studied apple genotypes, and the only detected putative QTL on linkage group 11 of'Fiesta' apples was not stable for the different replications of the experiment. This lack of quantifiable resistance may be partly explained by environmental conditions related to aphid development in sleeve cages.     

Molecular interactions between rosy apple aphids, Dysaphis plantaginea, and resistant and susceptible cultivars of its primary host Malus domestica

The use of crop varieties resistant or tolerant to insect pests or other stress factors is one approach in non‐chemical crop‐protection. Knowledge of the biochemical and molecular background of insect–plant interactions is a prerequisite for optimizing breeding for resistance. However, the resistance genes involved in plant–aphid interactions have so far only been identified and characterized in very few plant species. Our work aims to elucidate the molecular and biochemical mechanisms involved in resistance of apple trees, Malus domestica L. (Rosaceae), against its primary aphid pest, the rosy apple aphid, Dysaphis plantaginea (Passerini) (Homoptera: Aphididae), which is considered a serious economic pest of apple. Gene expression in both resistant and susceptible apple cultivars after infestation with rosy apple aphids was investigated by employing the cDNA‐AFLP method (cDNA–Amplified Fragment Length Polymorphism). From approximately 12 500 cDNA fragments detected on polyacrylamide gels, 21 bands were apparently up‐ or down‐regulated only in the resistant cultivar ‘Florina’ after aphid infestation compared to the susceptible cultivar ‘Topaz’ and/or mechanically wounded or non‐infested leaves. These fragments were cloned, sequenced, and the pattern of gene expression for six fragments was subsequently verified by virtual Northern blots. Sequence comparisons of these fragments to GenBank accessions revealed homologies to already known genes, most of them isolated from Arabidopsis thaliana L. Among them, a putative RNase‐L‐inhibitor‐like protein, a pectinacetylesterase, an inositol‐phosphatase‐like protein, a precursor of the large chain of the ribulose‐1,5‐biphosphate‐carboxylase, and defence‐related genes such as a vacuolar H(+)‐ATPase subunit‐like protein and an ADP‐ribosylating enzyme were identified. The results are discussed in relation to a putative role of these genes in conferring aphid resistance in apple trees.     

Genetic Control of Contagious Asexuality in the Pea Aphid

Although evolutionary transitions from sexual to asexual reproduction are frequent in eukaryotes, the genetic bases of such shifts toward asexuality remain largely unknown. We addressed this issue in an aphid species where both sexual and obligate asexual lineages coexist in natural populations. These sexual and asexual lineages may occasionally interbreed because some asexual lineages maintain a residual production of males potentially able to mate with the females produced by sexual lineages. Hence, this species is an ideal model to study the genetic basis of the loss of sexual reproduction with quantitative genetic and population genomic approaches. Our analysis of the co-segregation of ∼300 molecular markers and reproductive phenotype in experimental crosses pinpointed an X-linked region controlling obligate asexuality, this state of character being recessive. A population genetic analysis (>400-marker genome scan) on wild sexual and asexual genotypes from geographically distant populations under divergent selection for reproductive strategies detected a strong signature of divergent selection in the genomic region identified by the experimental crosses. These population genetic data confirm the implication of the candidate region in the control of reproductive mode in wild populations originating from 700 km apart. Patterns of genetic differentiation along chromosomes suggest bidirectional gene flow between populations with distinct reproductive modes, supporting contagious asexuality as a prevailing route to permanent parthenogenesis in pea aphids. This genetic system provides new insights into the mechanisms of coexistence of sexual and asexual aphid lineages.     

The use of common elder Sambucus nigra to promote Aphidophagous syrphids in apple orchards

Elder shrubs (Sambucus nigra L.) were planted in an experimental apple orchard as bordering hedgerow with the objective of rearing syrphids (Diptera: Syrphidae) and consequently enhancing the biological control of the rosy apple aphid Dysaphis plantaginea (Passerini). The inoculative introduction of the specific elder aphid Aphis sambuci L. (Homoptera: Aphididae) in late March 2002 and again in 2003 led to the complete hedgerow turning into an early and productive alternative prey reservoir for indigenous syrphids. The species Scaeva pyrastri (L.), S. selenetica (Meigen), Syrphus ribesii (L.), S. vitripennis Meigen, Epistrophe eligans (Harris), E. nitidicollis (Meigen), Platycheirus scutatus (Meigen), Eupeodes corollae (F.), Meligramma triangulifera (Zetterstedt) and Episyrphus balteatus (De Geer) developed on the elder hedgerow during the spring 2002 taking advantage of the aphid infestation. The season 2003 was characterised by a quite different species abundance pattern as, contrary to the previous year when both monovoltine and polyvoltine species were equally represented, the monovoltine syrphids (Epistrophe spp.) represented the dominant group exploiting the elder aphids. A parallel analysis showed that the faeces of the first adult syrphids observed ovipositing on the elder hedgerow in spring contained digested pollen of mainly apple. Our observations indicate the possibility of establishing a local population of monovoltine syrphids in apple orchards by managing an aphid-infested elder hedgerow, without any additional pollen or nectar producing plants.     

渭北塬区不同龄苹果园土壤微生物空间分布特征

以渭北塬区塬面5、10、15和20龄苹果园为对象,距树干1.0、1.5、2.0 m处用土钻法分层采集0—30 cm土样,稀释平板法测定土壤微生物数量,平行测定土壤有机碳、总氮和总磷,分析不同龄果园土壤微生物的空间分布状况及其与土壤碳、氮、磷的关系。结果表明:土壤真菌数量随树龄增大而增加,龄间差异显著(P0.05);与对照农田相比,各龄果园放线菌数量及5龄和20龄果园细菌数量偏低,10龄和15龄果园细菌数量偏高;各龄果园土壤真菌、细菌和放线菌数量均随深度增加而减小。就相同深度土层而言,真菌数量随果园年限增大而增加,细菌则以10龄和15龄果园较多,同深度土层的放线菌龄间差异随深度增加而减小。在沿树干向外的径向水平方向上,真菌数量随果园年限的增加相应增多;10龄和15龄果园土壤细菌和放线菌高于5龄果园和20龄果园。5龄果园土壤总氮有沿树干向外、沿表层向下逐步降低的趋势,果园从5龄经10龄到15龄,其"高氮点"则逐步向外、向下移动。塬区果园土壤C∶N比偏低。     

Spatial and temporal genetic variability in French populations of the peach-potato aphid,Myzus persicae

The peach-potato aphid, Myzus persicae (Sulzer), has a complex reproductive mode in which some lineages reproduce by continuous parthenogenesis, whereas others reproduce sexually once a year. The climate is thought to act directly on the reproductive mode, because sexual eggs are the only form that can resist frost in cold regions. Sexual reproduction necessitates an obligatory host alternation that may result in long-distance dispersal. Here, we examined the genetic variability at seven microsatellite loci of populations of M. persicae in France, where both reproductive modes occur. We provide clear genetic evidence that the breeding system affects genotypic variability, as cyclically parthenogenetic aphids are far more variable than their obligately parthenogenetic counterparts. A temporal decrease in genetic variability and a temporal genetic differentiation effect suggest the existence of selective factors that play an important role in shaping the genetic structure of M. persicae populations. Lastly, differences in the population structure between reproductive modes suggest that the migration associated with the change of host during sexual reproduction lowers the level of population differentiation.     

Genetic architecture of sexual and asexual populations of the aphid Rhopalosiphum padi based on allozyme and microsatellite markers

Cyclical parthenogens, including aphids, are attractive models for comparing the genetic outcomes of sexual and asexual reproduction, which determine their respective evolutionary advantages. In this study, we examined how reproductive mode shapes genetic structure of sexual (cyclically parthenogenetic) and asexual (obligately parthenogenetic) populations of the aphid Rhopalosiphum padi by comparing microsatellite and allozyme data sets. Allozymes showed little polymorphism, confirming earlier studies with these markers. In contrast, microsatellite loci were highly polymorphic and showed patterns very discordant from allozyme loci. In particular, microsatellites revealed strong heterozygote excess in asexual populations, whereas allozymes showed heterozygote deficits. Various hypotheses are explored that could account for the conflicting results of these two types of genetic markers. A strong differentiation between reproductive modes was found with both types of markers. Microsatellites indicated that sexual populations have high allelic polymorphism and heterozygote deficits (possibly because of population subdivision, inbreeding or selection). Little geographical differentiation was found among sexual populations confirming the large dispersal ability of this aphid. In contrast, asexual populations showed less allelic polymorphism but high heterozygosity at most loci. Two alternative hypotheses are proposed to explain this heterozygosity excess: allele sequence divergence during long-term asexuality or hybrid origin of asexual lineages. Clonal diversity of asexual lineages of R. padi was substantial suggesting that they could have frozen genetic diversity from the pool of sexual lineages. Several widespread asexual genotypes were found to persist through time, as already seen in other aphid species, a feature seemingly consistent with the general-purpose genotype hypothesis.